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(Carnivora, Phocidae): Middle Miocene Seals

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(Carnivora, Phocidae): Middle Miocene Seals ivist stlin di leontologi e trtigrfi volume IIW noF Q ppF QPSEQSH xovemer PHIQ ps igyh yp pysv gyrysxei @gexsyeD rygsheiAX wshhvi wsygixi iev pyw ri xyrix eeir ssxe eF uyiu 8 vwex tF erwe eeivedX epril IPD PHIQY eptedX tune PSD PHIQ uey wordsX hoideD gystophorineD phyosteoslerosisD seE inusuli per l9inter fmigliF sl nuovo genere del wioene ondivide xul dimorphismD wioeneD widdle rmtinD rtethysD krineF numerosi rtteri on diverse speie eenti di gystophor e wirounE gD ed offre l prim opportunitÁdi studire il dimorfismo sessule delle estrtF hespite long history of phoid studiesD no fossil oss degli rti e delle mndiole nell ottofmigli gystophorineF sl memers of the ufmily gystophorine hve ever een desriedF dimorfismo sessule nelle oss postrnili e nelle mndiole eÁ mgE xew fossil mteril from the widdle rmtin @IIFPEIPFQ wA in the giormente evidente nei memri ttuli di gystophorine rispetto gli rtethyn fsin of krine llows emended dignoses nd redesripE ltri rppresentnti delle fohe in senso strettoF v9nlisi dei trtti nE tions to help lrify phylogeneti reltionships within the pmily hoE tomii h dimostrto he entrme le nuove speie sono piuÁprimitive e ideF efter ldisti nd morphologil nlyses of the mterilD new meglio dttte per l loomozione terrestre di qulsisi ltro rppreE genus @hyphoA ws eretedD with two new speies of extint fossil sentnte vivente di gystophorineF hypho ukriniD piuÁpiolD eÁ true sels @hypho ukrini nd hypho hpskiiAD elonging meglio dttt ll loomozione terrestre rispetto ll speie piuÁgrnE to the ufmily gystophorineF his new mteril shows exeptionl de F hpskiiF v9nlisi filogeneti suggerise he le fohe on IH iniE phyosteosleroti onesD whih is unommon for the fmily s sivi @hoineA sono piuÁprimitive di quelle on V @wonhineAD e he i wholeF he new wioene genus shres numerous hrters with seE wonhine sono piuÁprimitivi delle fohe on T inisivi @gystophoriE verl eent speies of gystophor nd wiroungD providing the first neAF uesti ritrovmenti indino he l ottofmigli gystophorine opportunity to study sexul dimorphism of lim ones nd mndiles inlude non solo elefnti mrini e fohe dl ppuioD m nhe le due in the ufmily gystophorineF exul dimorphism in postrnil nuove fohe phiosteoslerotihe del rmtino wedioF ones nd mndiles in living memers of gystophorine is more oE vious thn in other representtives of true selsF ixmintion of ntoE mil trits demonstrted tht oth new speies re more primitive nd etter dpted for terrestril loomotion thn ny living representtives sntrodution of gystophorineF he smller hypho ukrini is more dpted to terrestril loomotion thn its lrger reltiveD F hpskiiF hylogeE he study of fossil sels of the northern flk e th neti nlysis suggests tht sels with IH inisors @hoineA re more entury with region @pigF IeA egn in the mid IW primitive thn those with V @wonhineAD nd tht wonhine re work y iihwld @IVSHD IVSQA nd xordmnn more primitive thn sels with T inisors @gystophorineAF hese finE @IVTHAD nd ws ontinued y endrusov @IVWQAD elekE dings indite tht the ufmily gystophorine inludes not only seev @IWPRAD imionesu @IWPSAD wrovii nd yesu elephnt nd hooded selsD ut lso the two new widdle rmtin phyosteosleroti selsF @IWRIAD wvren @IWTHAD uirpihnikov @IWTIAD uoretsky @PHHIAD nd uoretsky nd qrigoresu @PHHPAF e diverse issuntoF xonostnte l lung stori dello studio dei foidiD mix of true sels @hoideA hs een reported in these non er mi stto desritto un memro dell ottofmigli gystophoE studiesD ut no memers of the ytriideF rineF xuovo mterile fossile dl rmtino wedio @IIFPEIPFQ wA del hespite the undne nd rod distriution of fino dell rtetide dell9 rin permette or di emendre dignosi e desrizioni in modo d iutre hirire i rpporti filogenetii ll9inE hoideD mny prolems persist in interpreting the terno dell fmigli hoideF sn seguito ll9nlisi morfologi e ldiE systemtis of this groupD with the txonomi history sti del mterile eÁstto istituito un nuovo genere @hyphoAD on of gystophor eing espeilly onfusing @uoretsky 8 due nuove speie di fohe in senso stretto estinte @hypho ukrini role PHHPAF qry @IVTTA nd rouessrt @IVWUA divided e hypho hpskiiAD pprtenenti ll ottofmigli gystophorineF uesto nuovo mterile mostr oss phiosteoslerotihe eezionliD hoide into three sufmiliesX hoineD wonhine vortory of ivolutionry fiologyD heprtment of entomyD gollege of wediineD rowrd niversity SPH tF xD shingtonD hg PHHSW @ eAF iEmilX ikoretIPQdolFomY sulmnFrhmtdhowrdFedu QPT uoretsky sFeF 8 hmt FtF pigF I E eA leogeogrphi skethE mp of the istern rtethys for the durtion of the rmE tin e @IPFQ ± WFQT wA with redued slinity nd strong endemism @mp modified ftersvnov et lFD PHHUAF fA qenerlized geogrphil mp of krine showing the prinE ipl fossil lolities of the genus hypho from widE dle wioene @rmtinA of krine mentioned in the textX IA uhomutovo illge @RUëIT9IIFR99x QVëHW9QTFI99iAF PA qnylozuovo @RUëHT9STFQ99x QUëQI9TSFV99iAF QA oloty flk @RUëPP9RTFW99 x QQëSV9PQFS99iFAF RA heltokmenk @RUëRI9QRFP99x QQëSH9HTFV99iAF SA rhnkut @RSëPH9RTFW99x QPëPW9PQFS99iAF TA zunlr @RSëIS9VHFI99x QTëPP9PUFH99iAF UA qritsev @RWëSV9HSFP99x PUëIH9HQ99iAF nd gystophorineF hile there hs never een univerE inisors to PGIY PA possession of n infltle nsl sY QA sl eptne of this lssifitionD it ws followed y nonEdifferentited shpe @homodontyA of the postnine mny susequent workers inluding ygnev @IWQSAD teethY RA shpe nd diretion of mstoid proessY nd SA qrsse @IWSSAD heffer@IWSVAD uing @IWTRA nd ghpsE shpe of the mxillF ellen @IVVHAD followed y impson kii @IWURAF ghrters supporting the existene of three @IWRSAD reted n dditionl sufmilyD voodontineD sufmilies minly derive from nlyses of rnilD denE for southern sels tht ghpskii nd previous workers tl nd pelge morphologyF sn prtiulrD omprehenE regrded s trie within wonhineF sive nlysis of suprgeneri systemtis is found in the wjor divergenes from the threeEsufmily studies of ghpskii @IWSSD IWTID IWUID IWURAF re deE sheme egn with the work of uing @IWTTA nd furns sried dignosti rnil hrters @numer of iniE nd py @IWUHAF ghpskii @IWURA lso proved tht uing9s sorsD shpe of mxillD form of nterior pltl forminD @IWTTA hypothesis of ressigning the genus gystophor nd some othersA supporting the seprtion of true sels from the ufmily gystophorine into the ufmily into three sufmiliesD whih he in turn divided into hoine nd the genus wiroung into the ufmily tries nd suEtriesF wonhine is untenleF wuizon @IWVPA epted the hereforeD sels with six inisors elong to the systemtis of uingD nd returned to dismissing the uE ufmily gystophorine @qill IVTTAD omprising the fmily gystophorine s wholeF he orretness of gener gystophor xilssonD IVPH @hooded selA nd ghpskii9s onept ws orroorted y oinette wiroung qryD IVPU @elephnt sel or se elephntAF nd tins @IWUHA nd olly @PHHVA in theiromprtive his lssifition is sed on severl synpomorphies studies of the pinniped lneusD emphsizing tht it is etween gystophor nd wiroungX IA redution of the indmissile to txonomilly seprte the hooded sel widdle wioene sels from the xorthern rtethys QPU nd the se elephntF vterD this point of view ws lso teril @s for todyD only IH known fossil sel skulls hve supported y eninder @IWVHXUTAX FFF nlytil meE ever een found nd desriedA is tht the survivl rte thods of hromosome investigtions tully do not perE of fossil sel skulls is extremely low due to the pperE mit the seprtion of gener gystophor nd wiroungD thin thikness of skull onesF e similrondition pE nd this ontrsts with the onept of their seprte pers in otrideD ut fossil otride skull ones re txonomi sttus nd of inlusion of gystophor in muh denser nd hve higher survivl rteF hoine99F roweverD reent moleulr nd kryologil hue to the gret rrity nd usully unstisftory studies do not support this point of view nd ple preservtion of postrnilD nd espeilly rnilD fossil gystophor into the ufmily hoine nd wiroung remins of selsD ompred to terrestril rnivoresD this into the ufmily wonhine s seprte ldes @fiE remins one of the lest investigted nd most ompliE È nindEimonds 8 ussell IWWTY ernsson et lF PHHTY ted groups of lrge mmmlsF his study foused on pulton 8 troek PHIHAF iven moleulr iologists desriing previously unknown extint representtives onlude the need for reEevlution of pinniped txoE of gystophorine nd lrifying their reltionships with nomy @rigdon etF lF PHHUAF yurmorphologildt othersels of the pmily hoide nd its foursuE indites tht the new fossil sels from the widdle wioE fmilies @hoineD wonhineD gystophorineD nd ene @hyphoA hve omintion of ystophorine hevinophoineAF nd miroungini hrtersD reEopening the disussion emong xeogene mrine mmmlsD the remins one gin out the sufmilil reltionships of selsF of phoine sels @in omprison with monhines nd uing @IWTRAD furns nd py @IWUHAD grr nd erE ystophorinesA re reltively numerous in the widdle È ry @IWWRAD nd lter ernson et lF @PHHTA ssigned gysE rmtinEweotin deposits of the gentrl nd istern tophor to the rie gystophoriniD while wuenn nd rtethys @pigF IAF uh remins re found in the kE fell @IWWUA inluded gystophor in the rie hoiniF rineD woldviD omniD uzkhstnD lovkiD eusE iven todyD the ext txonomi reltionships of sels triD rungryD nd even in urkey @iihwld IVSHY within hoide re still ontroversil @yss IWWRY uorE xordmnn IVTHY elekseev IWPRY imionesu IWPSY etsky 8 qrigoresu PHHPY uoretsky 8 role PHHPAF wvren IWTHY qrigoresu IWUUY qrigoresu et lF rious sientists either seprte phoids intoX only IWVTY uoretsky IWVTD IWVUD D PHHIY uoretsky 8 qriE one sufmilyD hoine @yss IWVVY wuenn 8 fell goresu PHHPAF sn spite of the undne nd rod disE IWWUAY two sufmiliesD hoine nd wonhine triution of true selsD mny prolems persist in interE @furns 8
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